Sediment processing by the surgeonfish Ctenochaetus striatus at Moorea, French Polynesia

The sediment-feeding surgeonfish Ctenochaetus striatus was found to reduce the sizes of ingested carbonaceous particles. The mean sizes of particles in stomach samples of the surgeonfish were smaller than those from areas where the fish were observed feeding. The particle sizes in the rectal contents of the fish were smaller than those in the stomach samples. The sediment-feeding activities of this and related surgeonfishes are likely to have a local impact on the particle-size distributions of sediments in reef and lagoon habitats. The assimilation efficiencies of the fish were estimated by the ash-ratio method to be 20% for total organic matter and 37% for nitrogen.     

Patterns of migration between feeding and spawning sites in a coral reef surgeonfish

Many coral reef fishes exhibit regular localised migrations between feeding and spawning areas, but the factors affecting these migration patterns, such as the distance, frequency and spawning site fidelity are poorly understood. The aim of this study was to investigate the patterns of migration to spawning sites of the surgeonfish, Ctenochaetus striatus (Acanthuridae). We explored relationships amongst an individual’s size and sex, the distance and frequency it migrated from its feeding area to spawning sites, fidelity to particular spawning sites and the number of individuals that aggregated to spawn. In order to achieve this, 406 C. striatus were captured and tagged on inshore reefs in Kimbe Bay (5°30′S 150°6′E), New Britain, Papua New Guinea. Tagged individuals were consistently observed within spatially discrete but overlapping feeding areas (maximum diameter averaging <13 m). The mean distance migrated was 58 m (ranging from 2 to 291 m). No tagged individuals were witnessed spawning at more than one site. Whilst most individuals (n = 88) migrated to the spawning site that was closest to their feeding areas, those that migrated to sites further away (n = 9) always spawned at sites where the number of conspecifics aggregating was larger. Neither the size nor the sex of individuals limited migration distance. However, males migrated significantly more frequently than females (on average once every 2 days vs. once every 3 days), and migration frequency was positively correlated with size in females. Migration distance did not affect the frequency with which individuals spawned. Whether patterns of migration are determined by cost-benefit optimisation, tradition, or an alternative mechanism is unknown.     

Recruitment,loss and coexistence in a guild of territorial coral reef fishes

Summary Eupomacentrus apicalis, Plectroglyphidodon lacrymatus, and Pomacentrus wardi are territorial pomacentrid fishes which occupy contiguous individual territories within rubble patches on the shallow reef slope. Loss of residents, which is non-seasonal (except for juvenile Po. wardi), results in reallocation of space in rubble patches among the species. This reallocation is random in the sense that sites previously held by one species will not be more likely than any other sites to be reoccupied by that species or to be occupied by any other particular species. The results of a 38 month study of three neighbouring patches are used to determine patterns of recruitment, survivorship, and loss for each species. The rate of recruitment of fish is proportional to the area of the rubble patch, and is seasonal in at least one of the species. About half the recruits are juveniles and young adults from other sites. The others are newly settled from the plankton. Total space used in a rubble patch does not vary significantly during the 38 month period. Po. wardi recruits and is lost at higher rates than the other species and its survivorship is significantly lower. Juvenile Po. wardi are lost at a greater rate than are adults, and their loss rate varies seasonally. The other species are similar to one another in having low recruitment and loss rates, and correspondingly high survivorship. The data are used in a critical assessment of several competing hypotheses to explain the coexistence of these fish. The available data are most closely compatible with the chance allocation or lottery hypothesis, but a definitive conclusion is not possible, and is probably beyond current experimental techniques.     

A simple model of growth form-dependent recovery from disease in coral reef sponges, and implications for monitoring

For clonal organisms that can suffer high levels of partial mortality and still recover, the conditions that influence infection and development of disease (e.g., abiotic stressors, population density) may be very different from the conditions that influence recovery. Recovery from infectious disease may increase if an individual can mount a defense before infection spreads throughout its body. If pathogens spread within an organism from an initial infection point, growth form—in conjunction with size—can influence the amount of time before all the tissue is diseased, and recovery precluded. A simple model of pathogen progression within individual sponges predicts that species with massive growth forms will be most susceptible to being overwhelmed by pathogen infection, and branching species will be most likely to recover. These predictions may help to explain the seemingly contradictory observations that branching species had the greatest prevalence of disease, and massive species the greatest rate of loss, in a monitored coral reef community. Disease may be observed disproportionately frequently in the organisms that are most likely to recover, resulting in underestimation, by standard monitoring procedures, of the effect of disease on losses from the community.     

Recruitment and post-recruit immigration affect the local population size of coral reef fishes

 This study quantifies the contributions of larval recruitment and post-recruit (juvenile and adult) immigration to net increases in population size for 150 species of fishes found on ten isolated coral patches or ‘bommies’ (108–267 m²) within a typical reef of the Great Barrier Reef system. At least one third of the total number of recruits and immigrants to all bommies were post-recruit fishes, and movement between bommies in 136 species was detected at some time during the 22 month sampling period. The relative numbers of recruits and post-recruit immigrants per species varied widely within the assemblage, and between the replicate bommies. Populations of 95 species received both types of immigrants, 41 species had only post-recruit immigrants, and 14 species received only larval recruitment. In most species, recruitment occurred over the austral summer between October and February, while post-recruit movements occurred in both summer and winter. Rates of post-recruit immigration varied temporally within bommies, and pulses of post-recruits were less temporally concordant between bommies than pulses of recruits. This study is further evidence that post-settlement processes can have a significant effect on the local population size of reef fishes. Accepted: 30 January 1997     

Spatial variation in mechanical properties of coral reef substrate and implications for coral colony integrity

The physical structure of coral reefs plays a critical role as a barrier to storm waves and tsunamis and as a habitat for living reef-building and reef-associated organisms. However, the mechanical properties of reef substrate (i.e. the non-living benthos) are largely unknown, despite the fact that substrate properties may ultimately determine where organisms can persist. We used a geo-mechanical technique to measure substrate material density and strength over a reef hydrodynamic gradient. Contrary to expectation, we found a weak relationship between substrate strength and wave-induced water flow: flow rates decline sharply at the reef crest, whereas substrate properties are relatively constant over much of the reef before declining by almost an order of magnitude at the reef back. These gradients generate a novel hump-shaped pattern in resistance to mechanical disturbances for live corals, where colonies closer to the back reef are prone to dislodgement because of poorly cemented substrate. Our results help explain an intermediate zone of higher taxonomic and morphological diversity bounded by lower diversity exposed reef crest and unstable reef back zones.     

Effect of the surgeonfish <Emphasis Type="Italic">Ctenochaetus striatus</Emphasis> (Acanthuridae) on the processes of sediment transport and deposition on a coral reef in the Red Sea

Excessive sedimentation is a major threat to coral reefs. It can damage or kill reef-building corals and can prevent the successful settlement of their planktonic larvae. The surgeonfish Ctenochaetus striatus feeds on rocky surfaces by sweeping loose material into its mouth with its flexible, broom-like teeth. In addition, it grasps and removes hard substrates with the aid of its special palate structure. It then transports sediment matter off the reef by defecating the ingested material outside the rocky zone of the reef. We analyzed 150 feces samples of six individuals, differentiating between (1) ingested by sweeping and (2) ingested by scraping, and compared their content with inorganic land-derived and marine sediments trapped at the feeding area. Projections based on fish densities, defecation rates, and quantities as well as composition of sediments collected by traps on the same reef site suggest that C. striatus removes at least 18% of the inorganic sediment sinking onto the reef crest. The eroded share in the exported matter is about 13%. This finding points to a hitherto not verified role of C. striatus as a reef sweeper and reef scraper, whereby the first function is by far dominating.     

Cluster dynamics in Maristentor dinoferus,a gregarious benthic ciliate with zooxanthellae and a hypericin-like pigment,in relation to biofilm grazing by the fish Ctenochaetus striatus

The large symbiotic ciliate Maristentor dinoferus (Heterotrichida: Maristentoridae) has an abundant, potentially toxic pigment that makes the cells look black, and it forms clusters making it easy to see in the field. In situ observations of vast Maristentor populations during three bloom years provided insights into the behavioral ecology of the Maristentor holobiont and circumstantial evidence for feeding deterrence. Maristentor migrated onto limestone tiles that were also suitable for development of biofilm, which was consumed by the fish Ctenochaetus striatus. We photodocumented several aspects of Maristentor movement in situ, including a diurnal rhythm of dispersal and re-clustering in the morning, a period of “cocktail party dynamics” that resulted in larger, fewer clusters over the afternoon, and responses to two different disturbances: (1) rapid dispersal–re-clustering when tiles were moved; and (2) swimming off the surface and photoaccumulating on the brightest and highest part of a container when Maristentor was enclosed. Although Maristentor is considered benthic, its abilities to swim rapidly, to photoaccumulate, and to form clusters hanging from a water surface hint that they may also exist in the hyponeuston. Although interaction between Maristentor and Ctenochaetus may not be common, it indicated ways in which the behavior of Maristentor could help defend it from grazing. First, during the morning diaspora and re-clustering, which coincided with a period when Ctenochaetus was not feeding, Maristentor tended to accumulate on thinner biofilm, including edges and previously-grazed areas. Secondly, analysis of bite marks suggested that this fish tended not to select thin biofilm and that, despite rapid feeding, it tended to avoid larger Maristentor clusters.     

Interactions between coral restoration and fish assemblages: implications for reef management

Corals create complex reef structures that provide both habitat and food for many fish species. Because of numerous natural and anthropogenic threats, many coral reefs are currently being degraded, endangering the fish assemblages they support. Coral reef restoration, an active ecological management tool, may help reverse some of the current trends in reef degradation through the transplantation of stony corals. Although restoration techniques have been extensively reviewed in relation to coral survival, our understanding of the effects of adding live coral cover and complexity on fishes is in its infancy with a lack of scientifically validated research. This study reviews the limited data on reef restoration and fish assemblages, and complements this with the more extensive understanding of complex interactions between natural reefs and fishes and how this might inform restoration efforts. It also discusses which key fish species or functional groups may promote, facilitate or inhibit restoration efforts and, in turn, how restoration efforts can be optimised to enhance coral fish assemblages. By highlighting critical knowledge gaps in relation to fishes and restoration interactions, the study aims to stimulate research into the role of reef fishes in restoration projects. A greater understanding of the functional roles of reef fishes would also help inform whether restoration projects can return fish assemblages to their natural compositions or whether alternative species compositions develop, and over what timeframe. Although alleviation of local and global reef stressors remains a priority, reef restoration is an important tool; an increased understanding of the interactions between replanted corals and the fishes they support is critical for ensuring its success for people and nature.     

Near-surface enrichment of zooplankton over a shallow back reef: implications for coral reef food webs

Zooplankton were 3–8 times more abundant during the day near the surface than elsewhere in the water column over a 1–2.4 m deep back reef in Moorea, French Polynesia. Zooplankton were also significantly more abundant near the surface at night although gradients were most pronounced under moonlight. Zooplankton in a unidirectional current became concentrated near the surface within 2 m of departing a well-mixed trough immediately behind the reef crest, indicating that upward swimming behavior, rather than near-bottom depletion by reef planktivores, was the proximal cause of these gradients. Zooplankton were highly enriched near the surface before and after a full lunar eclipse but distributed evenly throughout the water column during the eclipse itself supporting light as a proximal cue for the upward swimming behavior of many taxa. This is the first investigation of the vertical distribution of zooplankton over a shallow back reef typical of island barrier reef systems common around the world. Previous studies on deeper fringing reefs found zooplankton depletion near the bottom but no enrichment aloft. In Moorea, where seawater is continuously recirculated out the lagoon and back across the reef crest onto the back reef, selection for upward swimming behavior may be especially strong, because the surface serves both as a refuge from predation and an optimum location for retention within the reef system. Planktivorous fish and corals that can forage or grow even marginally higher in the water column might have a substantial competitive advantage over those nearer the bottom on shallow reefs. Zooplankton abundance varied more over a few tens of centimeters vertical distance than it did between seasons or even between day and night indicating that great care must be taken to accurately assess the availability of zooplankton as food on shallow reefs.     

Radiographic studies of reef coral exoskeletons: Rates and patterns of coral growth

X-radiographs were made of vertical slices through the centers of 47 hermatypic coral colonies collected at Eniwetok Atoll, Marshall Islands. The image thus obtained are useful for the study of colony geometry, development, and response to damage.Comparison of radioactive inclusions of known age with previously reported cyclic skeletal density variations normal to the axis of growth confirms the annual nature of the density banding. Growth rates based on density bands and radioactivity inclusions are calculated for all 47 specimens, and measurements of the individual ‘growth bands’ are presented for 25 of them.Bulk densities measured by X-ray transmission ranged from 1.0 to 2.2 g/cm³, with an average range of 1.3–1.6 g/cm³. Intra-specimen skeletal densities typically vary by 10–30%; the period of high density skeletal deposition appears to coincide with the season of higher rainfall and warmer surface water at Eniwetok. Pigment residues left by boring algae are more commonly found in low density portions of the skeletons, but this distribution is believed to result from rather than cause the variations in the density of the deposited aragonite.Linear growth rates for the same specimen vary by factors of two or more from year to year, but the 25 specimens studied did not show a common pattern in the linear growth rate. Other than showing some general trends in growth as a function of species and depth, linear growth rates do not appear to be a particularly informative parameter.The density and growth rate variations are important factors in the measurement of coral growth and metabolism, and to the study of environmental controls of coral growth.     

Effects of herbivory,nutrients, and reef protection on algal proliferation and coral growth on a tropical reef

Maintaining coral reef resilience against increasing anthropogenic disturbance is critical for effective reef management. Resilience is partially determined by how processes, such as herbivory and nutrient supply, affect coral recovery versus macroalgal proliferation following disturbances. However, the relative effects of herbivory versus nutrient enrichment on algal proliferation remain debated. Here, we manipulated herbivory and nutrients on a coral-dominated reef protected from fishing, and on an adjacent macroalgal-dominated reef subject to fishing and riverine discharge, over 152 days. On both reefs, herbivore exclusion increased total and upright macroalgal cover by 9-46 times, upright macroalgal biomass by 23-84 times, and cyanobacteria cover by 0-27 times, but decreased cover of encrusting coralline algae by 46-100% and short turf algae by 14-39%. In contrast, nutrient enrichment had no effect on algal proliferation, but suppressed cover of total macroalgae (by 33-42%) and cyanobacteria (by 71% on the protected reef) when herbivores were excluded. Herbivore exclusion, but not nutrient enrichment, also increased sediment accumulation, suggesting a strong link between herbivory, macroalgal growth, and sediment retention. Growth rates of the corals Porites cylindrica and Acropora millepora were 30-35% greater on the protected versus fished reef, but nutrient and herbivore manipulations within a site did not affect coral growth. Cumulatively, these data suggest that herbivory rather than eutrophication plays the dominant role in mediating macroalgal proliferation, that macroalgae trap sediments that may further suppress herbivory and enhance macroalgal dominance, and that corals are relatively resistant to damage from some macroalgae but are significantly impacted by ambient reef condition.     

Habitat dynamics,marine reserve status,and the decline and recovery of coral reef fish communities

Severe climatic disturbance events often have major impacts on coral reef communities, generating cycles of decline and recovery, and in some extreme cases, community‐level phase shifts from coral‐ to algal‐dominated states. Benthic habitat changes directly affect reef fish communities, with low coral cover usually associated with low fish diversity and abundance. No‐take marine reserves (NTRs) are widely advocated for conserving biodiversity and enhancing the sustainability of exploited fish populations. Numerous studies have documented positive ecological and socio‐economic benefits of NTRs; however, the ability of NTRs to ameliorate the effects of acute disturbances on coral reefs has seldom been investigated. Here, we test these factors by tracking the dynamics of benthic and fish communities, including the important fishery species, coral trout (Plectropomus spp.), over 8 years in both NTRs and fished areas in the Keppel Island group, Great Barrier Reef, Australia. Two major disturbances impacted the reefs during the monitoring period, a coral bleaching event in 2006 and a freshwater flood plume in 2011. Both disturbances generated significant declines in coral cover and habitat complexity, with subsequent declines in fish abundance and diversity, and pronounced shifts in fish assemblage structure. Coral trout density also declined in response to the loss of live coral, however, the approximately 2:1 density ratio between NTRs and fished zones was maintained over time. The only post‐disturbance refuges for coral trout spawning stocks were within the NTRs that escaped the worst effects of the disturbances. Although NTRs had little discernible effect on the temporal dynamics of benthic or fish communities, it was evident that the post‐disturbance refuges for coral trout spawning stocks within some NTRs may be critically important to regional‐scale population persistence and recovery.     

Survival dynamics of scleractinian coral larvae and implications for dispersal

Survival of pelagic marine larvae is an important determinant of dispersal potential. Despite this, few estimates of larval survival are available. For scleractinian corals, few studies of larval survival are long enough to provide accurate estimates of longevity. Moreover, changes in mortality rates during larval life, expected on theoretical grounds, have implications for the degree of connectivity among reefs and have not been quantified for any coral species. This study quantified the survival of larvae from five broadcast-spawning scleractinian corals (Acropora latistella, Favia pallida, Pectinia paeonia, Goniastrea aspera, and Montastraea magnistellata) to estimate larval longevity, and to test for changes in mortality rates as larvae age. Maximum lifespans ranged from 195 to 244 d. These longevities substantially exceed those documented previously for coral larvae that lack zooxanthellae, and they exceed predictions based on metabolic rates prevailing early in larval life. In addition, larval mortality rates exhibited strong patterns of variation throughout the larval stage. Three periods were identified in four species: high initial rates of mortality; followed by a low, approximately constant rate of mortality; and finally, progressively increasing mortality after approximately 100 d. The lifetimes observed in this study suggest that the potential for long-distance dispersal may be substantially greater than previously thought. Indeed, detection of increasing mortality rates late in life suggests that energy reserves do not reach critically low levels until approximately 100 d after spawning. Conversely, increased mortality rates early in life decrease the likelihood that larvae transported away from their natal reef will survive to reach nearby reefs, and thus decrease connectivity at regional scales. These results show how variation in larval survivorship with age may help to explain the seeming paradox of high genetic structure at metapopulation scales, coupled with the maintenance of extensive geographic ranges observed in many coral species. Communicated by Environment Editor Prof. van Woesik.     

Patch dynamics of coral reef macroalgae under chronic and acute disturbance

The patch dynamics (colonisation rate, growth rate, and extinction rate) are quantified for two dominant species of macroalgae on a Caribbean forereef in Belize: Lobophora variegata (Lamouroux) and Dictyota pulchella (Hörnig and Schnetter). Measurements were taken on time scales of days, weeks, months, and years during which three hurricanes occurred. All patches were followed on naturally occurring ramets of dead Montastraea annularis. The first hurricane (Mitch) caused massive coral mortality and liberated space for algal colonisation. The cover of Lobophora increased throughout the study and herbivores did not appear to limit its cover within a 4 year time frame. In contrast, the cover of D. pulchella fluctuated greatly and showed no net increase, despite an increase in parrotfish biomass and settlement space. Variation in the overall percent cover of an alga is not indicative of the underlying patch dynamics. The steady rise in the cover of Lobophora took place despite a high turnover of patches (12–60% of patches per year). The patch dynamics of Dictyota were slower (7–20%), but a greater patch density and threefold higher lateral growth rate led to greater fluctuations in total cover. The dynamics of algal patches are size-specific such that larger patches are less likely to become extinct during hurricanes.     

Larval dispersal and movement patterns of coral reef fishes,and implications for marine reserve network design

Well‐designed and effectively managed networks of marine reserves can be effective tools for both fisheries management and biodiversity conservation. Connectivity, the demographic linking of local populations through the dispersal of individuals as larvae, juveniles or adults, is a key ecological factor to consider in marine reserve design, since it has important implications for the persistence of metapopulations and their recovery from disturbance. For marine reserves to protect biodiversity and enhance populations of species in fished areas, they must be able to sustain focal species (particularly fishery species) within their boundaries, and be spaced such that they can function as mutually replenishing networks whilst providing recruitment subsidies to fished areas. Thus the configuration (size, spacing and location) of individual reserves within a network should be informed by larval dispersal and movement patterns of the species for which protection is required. In the past, empirical data regarding larval dispersal and movement patterns of adults and juveniles of many tropical marine species have been unavailable or inaccessible to practitioners responsible for marine reserve design. Recent empirical studies using new technologies have also provided fresh insights into movement patterns of many species and redefined our understanding of connectivity among populations through larval dispersal. Our review of movement patterns of 34 families (210 species) of coral reef fishes demonstrates that movement patterns (home ranges, ontogenetic shifts and spawning migrations) vary among and within species, and are influenced by a range of factors (e.g. size, sex, behaviour, density, habitat characteristics, season, tide and time of day). Some species move <0.1–0.5 km (e.g. damselfishes, butterflyfishes and angelfishes), <0.5–3 km (e.g. most parrotfishes, goatfishes and surgeonfishes) or 3–10 km (e.g. large parrotfishes and wrasses), while others move tens to hundreds (e.g. some groupers, emperors, snappers and jacks) or thousands of kilometres (e.g. some sharks and tuna). Larval dispersal distances tend to be <5–15 km, and self‐recruitment is common. Synthesising this information allows us, for the first time, to provide species, specific advice on the size, spacing and location of marine reserves in tropical marine ecosystems to maximise benefits for conservation and fisheries management for a range of taxa. We recommend that: (i) marine reserves should be more than twice the size of the home range of focal species (in all directions), thus marine reserves of various sizes will be required depending on which species require protection, how far they move, and if other effective protection is in place outside reserves; (ii) reserve spacing should be <15 km, with smaller reserves spaced more closely; and (iii) marine reserves should include habitats that are critical to the life history of focal species (e.g. home ranges, nursery grounds, migration corridors and spawning aggregations), and be located to accommodate movement patterns among these. We also provide practical advice for practitioners on how to use this information to design, evaluate and monitor the effectiveness of marine reserve networks within broader ecological, socioeconomic and management contexts.     

Survival dynamics of reef coral larvae with special consideration of larval size and the genus Acropora

Temporal dynamics of larval survival were examined in vitro in four broadcast-spawning reef coral species, Acropora hyacinthus, A. japonica, A. solitaryensis, and Goniastrea pectinata. Larval size was treated as an important characteristic that may relate to larval lifespan. Two patterns were observed in larval survival dynamics between the three Acropora species (mean initial larval size; 0.05-0.08 mm(3)) and G. pectinata (0.02 mm(3)), based on the timing of a sharp drop in larval survival rates (ca. > 50% reduction over a 1-2 week period). Consequently, the majority of larvae of the three Acropora species had a lifespan of less than 2-3 weeks, whereas those of G. pectinata were extended a further 2-3 weeks despite the smaller larval size. No significant relationship was detected between the initial larval size and larval lifespan in any of the four reef coral species. These results suggest that (1) larval dispersal of spawning Acropora species may be on a more local scale than that of G. pectinata and most other reef coral species previously reported, and (2) larval size is not a good estimator of larval lifespan in reef coral species.     

Composition and sources of near reef zooplankton on a Jamaican forereef along with implications for coral feeding

Nocturnal near-reef zooplankton from the forereef of Discovery Bay, Jamaica, were sampled during winter and summer 1994 using a diver-operated plankton pump with an intake head positioned within centimeters of benthic zooplanktivores. The pump collected zooplankton not effectively sampled by conventional net tows or demersal traps. We found consistently greater densities of zooplankton than did earlier studies that used other sampling methods in similar locations. There was no significant difference between winter (3491±578 m^–3) and summer (2853±293 m^–3) zooplankton densities. Both oceanic- and reef-associated forms were found at temporal and spatial scales relevant to benthic suspension feeders. Copepods were always the most abundant group, averaging 89% of the total zooplankton, and most were not of demersal origin. The cyclopoids, Oithona spp., were the numerically dominant organisms, with an average density of 1684±260 m^–3. Other zooplankton (e.g., shrimp larvae, crab larvae, polychaetes, chaetognaths, amphipods, and isopods) were highly variable and much less abundant. Near-reef zooplankton abundances were high throughout the night sampling period, not just after sunset and before sunrise as previously described. Mean biomass was 4.5 mg C m^–3, with values ranging from 1.0 to 15.6 mg C m^–3. This work has important implications for evaluating which zooplankton types are available to benthic suspension feeders, including corals.